Long-Term Changes in the Avifauna of Barro Colorado Island, Panama, a Tropical Forest Isolate

Since its isolation from the mainland more than 85 years ago, 65 bird species have disappeared from Barro Colorado Island, Panama. Because of these extinctions the island is often used as a model for the study of faunal relaxation, or loss of species through time. The most recent survey of the island's bird community was completed in 1970. Between June 1994 and May 1996 I surveyed the island and observed 218 species, including 5 species never before recorded. Three relatively sedentary species have experienced severe population declines since Willis's 1970 survey and may be on the brink of disappearing from the island. Willis estimated 500 Slate-colored Grosbeaks ( Pitylus grossus ), whereas I found only two pairs of this conspicuous midstory-dwelling finch. Two cotingas—Speckled Mourner ( Laniocera rufescens ) and Rufous Piha ( Lipaugus unirufus )—have declined by 85–95%. I did not locate any individuals of 36 other species that were present during the 1970s. Most (21) of these are uncommon aquatic or nocturnal species, which I may have missed during my surveys, or are rare to uncommon edge species that probably move frequently to and from the island. Seven species, however, are primarily inhabitants of second-growth forest and have been lost probably because of continuing successional maturation of the forest, including changes in land use around the laboratory clearing. Seven forest-dwelling species disappeared during the 1970s and have not recolonized. I detected only one sedentary forest-dwelling species, Great Currasow ( Crax rubra ), that previously had been reported as missing from the island. The nearly complete lack of recolonizations by such forest-interior species suggests that local extinction from tropical forest isolates may be extremely persistent. Tropical forest reserves as small as Barro Colorado Island (1600 ha) may not preserve high levels of regional avian diversity over long periods of time.     

The phenology of a Neotropical ant assemblage: evidence for continuous and overlapping reproduction

Reproductive phenologies reflect the interaction between the mating system of a taxon and the local environment. Ant colonies reproduce and disperse via the flights of winged alates. Few data exist on the reproductive phenologies of ant assemblages. Here we analyze the reproductive phenologies of 81 common ant species from 23,182 individuals collected over 3 years on Barro Colorado Island, Panama (BCI). Species ranged from highly synchronous to continuous fliers, but showed a median flight duration of at least 8 of 13 lunar months. In two statistical analyses (variance ratio test and Spearman rank correlations), 84% (16 of 19) of ant genera had species trending toward positively associated phenologies, more than expected by chance (P<0.00036 by a binomial test). Thus, there was little evidence for the hypothesis that competition for limiting resources staggers congeneric flights and ultimately promotes reproductive isolation. On the contrary, the timing of reproduction, and its synchrony, tended to be conserved within genera and subfamilies. These results closely match phenological studies of plant assemblages. The continuous reproduction and small colony size of many species in this study suggest that the female calling syndrome, a poorly documented mating system in ants, may be common on BCI.     

Echolocation beam shape in emballonurid bats, Saccopteryx bilineata and Cormura brevirostris

The shape of the sonar beam plays a crucial role in how echolocating bats perceive their surroundings. Signal design may thus be adapted to optimize beam shape to a given context. Studies suggest that this is indeed true for vespertilionid bats, but little is known from the remaining 16 families of echolocating bats. We investigated the echolocation beam shape of two species of emballonurid bats, Cormura brevirostris and Saccopteryx bilineata, while they navigated a large outdoor flight cage on Barro Colorado Island, Panama. C. brevirostris emitted more directional signals than did S. bilineata. The difference in directionality was due to a markedly different energy distribution in the calls. C. brevirostris emitted two call types, a multiharmonic shallowly frequency-modulated call and a multiharmonic sweep, both with most energy in the fifth harmonic around 68?kHz. S. bilineata emitted only one call type, multiharmonic shallowly frequency-modulated calls with most energy in the second harmonic (~46?kHz). When comparing same harmonic number, the directionality of the calls of the two bat species was nearly identical. However, the difference in energy distribution in the calls made the signals emitted by C. brevirostris more directional overall than those emitted by S. bilineata. We hypothesize that the upward shift in frequency exhibited by C. brevirostris serves to increase directionality, in order to generate a less cluttered auditory scene. The study indicates that emballonurid bats are forced to adjust their relative harmonic energy instead of adjusting the fundamental frequency, as the vespertilionids do, presumably due to a less flexible sound production.     

Directional changes in the species composition of a tropical forest

Long-term studies have revealed that the structure and dynamics of many tropical forests are changing, but the causes and consequences of these changes remain debated. To learn more about the forces driving changes within tropical forests, we investigated shifts in tree species composition over the past 25 years within the 50-ha Forest Dynamics Plot on Barro Colorado Island (BCI), Panama, and examined how observed patterns relate to predictions of (1) random population fluctuations, (2) carbon fertilization, (3) succession from past disturbance, (4) recovery from an extreme El Ni?o drought at the start of the study period, and (5) long-term climate change. We found that there have been consistent and directional changes in the tree species composition. These shifts have led to increased relative representations of drought-tolerant species as determined by the species' occurrence both across a gradient of soil moisture within BCI and across a wider precipitation gradient from a dry forest near the Pacific coast of Panama to a wet forest near its Caribbean coast. These nonrandom changes cannot be explained by stochastic fluctuations or carbon fertilization. They may be the legacy of the El Ni?o drought, or alternatively, potentially reflect increased aridity due to long-term climate change. By investigating compositional changes, we increased not only our understanding of the ecology of tropical forests and their responses to large-scale disturbances, but also our ability to predict how future global change will impact some of the critical services provided by these important ecosystems.     

Avian Survival Rates and the Extinction Process on Barro Colorado Island, Panama

Abstract: Selective extinction following isolation of habitat patches may be due to biogeographical (e.g., island size or isolation) and ecological (species natural histories, interspecifc interactions) factors, or their interactions. Among the demographic and life history attributes commonly associated with high extinction probability are small populations, large size of individuals, and population variability. Long-term capture-recapture data from forest habitat in central Panama permit an examination of the association between mainland survival rates and extinction on a nearby land-bridge island Species of birds that no longer occur on Barro Colorado Island (BCI), Panama, have, on average, lower survival rates on the adjacent mainland than species that have persisted on BCI. Moreover, of the species that no longer occur on BCI, those with lower mainland survival rates generally disappeared earlier from the island. My analysis provides little evidence of a relationship between extinction and population size. Recolonization of BCI from the adjacent mainland by the forest undergrowth species studied here is unlikely. Reduced reproductive success on BCI combined with naturally low adult survival rates seems to be responsible for these BCI extinctions. High nest predation and/or altered landscape dynamics are probable agents in the low reproductive success. The methods used here could be employed in other circumstances to identify fragmentation-sensitive species.     

Quantifying Rarity, Losses, and Risks for Native Fishes of the Lower Colorado River Basin: Implications for Conservation Listing

Abstract:  We examined spatial distributions of fishes native to the lower basin of the Colorado River (25 species) at three scales to determine percent decline from historical distributions based on a regional biodiversity database. We cumulated records from 1843 to 1980 to develop a "historical distribution" for each species and used those occurrences recorded from 1981 to 1998 as "modern" records. We then contrasted historical and modern distributions to (1) quantify losses in spatial distribution; (2) determine how strongly these losses and fragmentation patterns corresponded to the perceived risk of extinction of each species, as represented by its status under the IUCN Red List of Endangered Species; and (3) update extinction risk rankings for 15 fishes endemic to the lower Colorado Basin according to the IUCN criteria. Based on presence and absence data, fish fauna of the lower Colorado Basin have suffered massive distributional losses. On average, ranges of extant species have diminished more than 45% relative to their historical distribution, and 35% of species have lost 50% or more of their occurrences. We provide nine new IUCN rankings and six updates to reflect more accurately the heightened imperilment of these species. Based on our new rankings, 7 of the 15 lower Colorado Basin endemics are critically endangered, 1 is endangered, 2 are vulnerable, and 1 is already extinct. We categorize the remaining 2 endemics as lower risk. This work demonstrates the utility of matching quantitative spatial metrics such as the scale-area slope statistic to extinction risk criteria for species whose persistence is strongly influenced by spatial distribution.     

Compositional and functional stability of arthropod communities in the face of ant invasions

There is a general consensus that the diversity of a biotic community can have an influence on its stability, but the strength, ubiquity, and relative importance of this effect is less clear. In the context of biological invasions, diversity has usually been studied in terms of its effect on a community's invasibility, but diversity may also influence stability by affecting the magnitude of compositional or functional changes experienced by a community upon invasion. We examined the impacts of invasive ants on arthropod communities at five natural area sites in the Hawaiian Islands, and assessed whether differences among sites in community diversity and density variables were related to measures of stability. Ant invasion was usually associated with significant changes in overall community composition, as measured by Bray-Curtis distances, particularly among endemic subsets of the communities. Changes in mean species richness were also strong at three of the five sites. Among sites, diversity was negatively related to stability as measured by resistance to overall compositional change, but this effect could not be separated from the strong negative effect of invasive ant density on compositional stability. When compositional stability was measured as proportional change in richness, the best predictor of stability among endemic community subsets was endemic richness, with richer communities losing proportionately more species than species-poor communities. This effect was highly significant even after controlling for differences in invasive ant density and suggested that communities that had already lost many endemic species were resistant to further species loss upon ant invasion, while more intact communities remained vulnerable to species loss. Communities underwent strong but idiosyncratic functional shifts in association with ant invasion, both in terms of trophic structure and total arthropod biomass. There were no apparent relationships, however, between functional stability and community diversity or density measures. Instead, invasive ant density was the best among-site predictor of the magnitude of functional change. Overall, diversity appeared to be a poor predictor of stability in the face of ant invasion in these communities, possibly because any actual diversity effects were overshadowed by community-specific factors and variation in the magnitude of the ant-mediated perturbation.     

Food quality, competition, and parasitism influence feeding preference in a neotropical lepidopteran

We surveyed Lepidoptera found on 11 species of Inga (Fabaceae:Mimosoideae) co-existing on Barro Colorado Island, Panama, to evaluate factors influencing diet choice. Of the 47 species of caterpillars (747 individuals) recorded, each fed on a distinct set of Inga. In the field, 96% of the individuals were found on young leaves. Growth rates of caterpillars that were fed leaves in the laboratory were 60% higher on young leaves compared to mature leaves. When caterpillars were fed leaves of nonhost Inga, they grew more slowly. These data provide support for a link between preference and performance. However, among hosts on which larvae normally occurred, faster growth rates were not associated with greater host electivity (the proportion of larvae found on each host species in the field, corrected for host abundance). Growth rates on normal hosts were positively correlated with leaf expansion rates of the host, and fast expansion was associated with leaves with higher nutritional content. Detailed studies on a gelechiid leaf roller, the species with the largest diet breadth, allowed us to assess the importance of factors other than growth that could influence diet electivity. This species showed a 1.7-fold difference in growth rate among Inga hosts and faster growth on species with fast-expanding leaves. However, there was no correlation between caterpillar growth rate and abundance on different host species. Instead, abundance of the gelechiid on each Inga species was significantly correlated with the temporal predictability of food (synchrony of leaf flushing) and was negatively correlated with competition (amount of leaf area removed by species other than the gelechiid). Although rates of parasitism were high (23-43%), there were no differences among hosts. Parasitism was also not related to measures of escape, such as growth rates of caterpillars, leaf expansion rates, and synchrony of leaf production. Together, food availability, parasitism, and competition explained 84% of the variation in host preference by the gelechiid. We suggest that these ecological interactions may be particularly important in determining diet choice initially and that preferences may be reinforced by subsequent divergence in host chemistry and/or the herbivore's ability to tolerate the secondary metabolites.     

Life history trade-offs in tropical trees and lianas

It has been hypothesized that tropical trees partition forest light environments through a life history trade-off between juvenile growth and survival; however, the generality of this trade-off across life stages and functional groups has been questioned. We quantified trade-offs between growth and survival for trees and lianas on Barro Colorado Island (BCI), Panama using first-year seedlings of 22 liana and 31 tree species and saplings (10 mm < dbh < 39 mm) of 30 tree species. Lianas showed trade-offs similar to those of trees, with both groups exhibiting broadly overlapping ranges in survival and relative growth rates as seedlings. Life history strategies at the seedling stage were highly correlated with those at the sapling stage among tree species, with all species showing an increase in survival with size. Only one of 30 tree species demonstrated a statistically significant ontogenetic shift, having a relatively lower survival rate at the sapling stage than expected. Our results indicate that similar life history trade-offs apply across two functional groups (lianas and trees), and that life history strategies are largely conserved across seedling and sapling life-stages for most tropical tree species.     

Biodiversity Loss in Latin American Coffee Landscapes: Review of the Evidence on Ants,Birds, and Trees

Abstract: Studies have documented biodiversity losses due to intensification of coffee management (reduction in canopy richness and complexity). Nevertheless, questions remain regarding relative sensitivity of different taxa, habitat specialists, and functional groups, and whether implications for biodiversity conservation vary across regions. We quantitatively reviewed data from ant, bird, and tree biodiversity studies in coffee agroecosystems to address the following questions: Does species richness decline with intensification or with individual vegetation characteristics? Are there significant losses of species richness in coffee‐management systems compared with forests? Is species loss greater for forest species or for particular functional groups? and Are ants or birds more strongly affected by intensification? Across studies, ant and bird richness declined with management intensification and with changes in vegetation. Species richness of all ants and birds and of forest ant and bird species was lower in most coffee agroecosystems than in forests, but rustic coffee (grown under native forest canopies) had equal or greater ant and bird richness than nearby forests. Sun coffee (grown without canopy trees) sustained the highest species losses, and species loss of forest ant, bird, and tree species increased with management intensity. Losses of ant and bird species were similar, although losses of forest ants were more drastic in rustic coffee. Richness of migratory birds and of birds that forage across vegetation strata was less affected by intensification than richness of resident, canopy, and understory bird species. Rustic farms protected more species than other coffee systems, and loss of species depended greatly on habitat specialization and functional traits. We recommend that forest be protected, rustic coffee be promoted, and intensive coffee farms be restored by augmenting native tree density and richness and allowing growth of epiphytes. We also recommend that future research focus on potential trade‐offs between biodiversity conservation and farmer livelihoods stemming from coffee production.     

Effects of Dam‐Induced Landscape Fragmentation on Amazonian Ant–Plant Mutualistic Networks

Mutualistic networks are critical to biological diversity maintenance; however, their structures and functionality may be threatened by a swiftly changing world. In the Amazon, the increasing number of dams poses a large threat to biological diversity because they greatly alter and fragment the surrounding landscape. Tight coevolutionary interactions typical of tropical forests, such as the ant–myrmecophyte mutualism, where the myrmecophyte plants provide domatia nesting space to their symbiotic ants, may be jeopardized by the landscape changes caused by dams. We analyzed 31 ant–myrmecophyte mutualistic networks in undisturbed and disturbed sites surrounding Balbina, the largest Central Amazonian dam. We tested how ant–myrmecophyte networks differ among dam‐induced islands, lake edges, and undisturbed forests in terms of species richness, composition, structure, and robustness (number of species remaining in the network after partner extinctions). We also tested how landscape configuration in terms of area, isolation, shape, and neighborhood alters the structure of the ant–myrmecophyte networks on islands. Ant–myrmecophytic networks were highly compartmentalized in undisturbed forests, and the compartments had few strongly connected mutualistic partners. In contrast, networks at lake edges and on islands were not compartmentalized and were negatively affected by island area and isolation in terms of species richness, density, and composition. Habitat loss and fragmentation led to coextinction cascades that contributed to the elimination of entire ant–plant compartments. Furthermore, many myrmecophytic plants in disturbed sites lost their mutualistic ant partners or were colonized by opportunistic, nonspecialized ants. Robustness of ant–myrmecophyte networks on islands was lower than robustness near lake edges and in undisturbed forest and was particularly susceptible to the extinction of plants. Beyond the immediate habitat loss caused by the building of large dams in Amazonia, persistent edge effects and habitat fragmentation associated with dams had large negative effects on animal–plant mutualistic networks. Efectos de la Fragmentación del Paisaje Inducida por Presas sobre Redes Mutualistas Hormiga‐Planta Amazónicas     

Group size and growth rate in hatchling green iguanas (Iguana iguana)

Summary The post-hatching growth and sleeping group sizes of green iguanas (Iguana iguana) were followed on Barro Colorado Island, Panama, for 3 successive years. The laboratory clearing was subdivided and surveyed routinely for iguanas, most intensively the six weeks following onset of hatching. A similar pattern of influx of iguanas into the clearing took place each May. Over the early weeks the size of groups generally increased. Animals sleeping in larger groups had significantly faster growth rates. Animals that disappeared from the clearing prior to July had larger group sizes prior to disappearance than those that remained. Growth rates measured in animals that remained declined markedly after June. Possible reasons for the social grouping of young iguanas are discussed.     

Compensatory responses to loss of warming-sensitive plant species

Climate warming-induced plant species loss is likely to be nonrandom and based on species-specific susceptibility to changing climate. We examined the ecological consequences of losing shallow-rooted forbs, a group of species we predict to be adversely affected by climate change based on their response to experimental warming. After three years of experimental species removal, tap-rooted forbs and grasses were able to fully compensate for the loss of shallow-rooted forbs with increased biomass production. Moreover, the remaining plant community yielded a larger biomass response to nitrogen addition when shallow-rooted forbs were removed, possibly because removal led to increased soil moisture. We conclude that, although shallow-rooted forbs share a common response to warming, their loss did not affect community-level biomass. However, the loss of shallow-rooted forbs could result in increased sensitivity to perturbations, such as changing nutrient availability. Our results demonstrate that realistic, nonrandom scenarios of species loss do not necessarily follow the general pattern of decreased productivity and dampened response to nitrogen addition with species loss that is predicted by theory and many experimental results. Further examinations of nonrandom species loss in other ecosystems are needed to further improve our understanding of the consequences of human-driven species loss.     

Neutral theory and the evolution of ecological equivalence

Since the publication of the unified neutral theory in 2001, there has been much discussion of the theory, pro and con. The hypothesis of ecological equivalence is the fundamental yet controversial idea behind neutral theory. Assuming trophically similar species are demographically alike (symmetric) on a per capita basis is only an approximation, but it is equivalent to asking: How many of the patterns of ecological communities are the result of species similarities, rather than of species differences? The strategy behind neutral theory is to see how far one can get with the simplification of assuming ecological equivalence before introducing more complexity. In another paper, I review the empirical evidence that led me to hypothesize ecological equivalence among many of the tree species in the species-rich tropical forest on Barro Colorado Island (BCI). In this paper, I develop a simple model for the evolution of ecological equivalence or niche convergence, using as an example evolution of the suite of life history traits characteristic of shade tolerant tropical tree species. Although the model is simple, the conclusions from it seem likely to be robust. I conclude that ecological equivalence for resource use are likely to evolve easily and often, especially in species-rich communities that are dispersal and recruitment limited. In the case of the BCI forest, tree species are strongly dispersal- and recruitment-limited, not only because of restricted seed dispersal, but also because of low recruitment success due to heavy losses of the seedling stages to predators and pathogens and other abiotic stresses such as drought. These factors and the high species richness of the community strongly reduce the potential for competitive exclusion of functionally equivalent or nearly equivalent species.     

Patchy distribution of zooplankton: Behavior,population assessment and sampling problems

Zooplankton and micronekton are often highly aggregated. Density in aggregations can reach 100 to more than 1000 times the average density of the population as estimated by net sampling. In order to assess true abundance of the animals and understand the significance of the aggregations, more information is needed on the behavior and population ecology of individual species and the species-specific attributes of discrete assemblages. We present information on patterns of intense aggregations of protozoans, platyhelminthes, scyphomedusae, copepods, mysids, and sergestids from our own observations. Characteristics behaviors of different species include feeding swarms of Noctiluca miliaris, behaviorally maintained swarms of planktonic flatworms, diel horizontal migration of Mastigias sp. swarms, three patterns of copepod swarms, bathymetric zonation of mysid schools, and seasonal migration and near bottom swarming of Sergia lucens. After the principles were demonstrated, we consider how to relate sampling, behavior, and population dynamics. We stress that the methodology is the result of the question, not the other way round. To tackle the problems associated with behavior in the open ocean, which we did not observe from the surface, it is necessary to sample in more than one way to extract data on differences in population-specific biology that alternative methods supply.     

Functional traits and the growth-mortality trade-off in tropical trees

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 < or = 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (H(max)) are key traits among forest trees. Seed mass, LMA, WD, and H(max) are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r2 < 0.1 for three and r2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for > 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.     

Biodiversity Conservation, Ecosystem Functioning, and Economic Incentives under Cocoa Agroforestry Intensification

Abstract:  World chocolate demand is expected to more than double by 2050. Decisions about how to meet this challenge will have profound effects on tropical rainforests and wild species in cocoa-producing countries. Cocoa, "the chocolate tree," is traditionally produced under a diverse and dense canopy of shade trees that provide habitat for a high diversity of organisms. The current trend to reduce or eliminate shade cover raises concerns about the potential loss of biodiversity. Nevertheless, few studies have assessed the ecological consequences and economic trade-offs under different management options in cocoa plantations. Here we describe the relationships between ant ecology (species richness, community composition, and abundance) and vegetation structure, ecosystem functions, and economic profitability under different land-use management systems in 17 traditional cocoa forest gardens in southern Cameroon. We calculated an index of profitability, based on the net annual income per hectare. We found significant differences associated with the different land-use management systems for species richness and abundance of ants and species richness and density of trees. Ant species richness was significantly higher in floristically and structurally diverse, low-intensity, old cocoa systems than in intensive young systems. Ant species richness was significantly related to tree species richness and density. We found no clear relationship between profitability and biodiversity. Nevertheless, we suggest that improving the income and livelihood of smallholder cocoa farmers will require economic incentives to discourage further intensification and ecologically detrimental loss of shade cover. Certification programs for shade-grown cocoa may provide socioeconomic incentives to slow intensification.     

Genetic Identification of Spotted Owls, Barred Owls, and Their Hybrids: Legal Implications of Hybrid Identity

Abstract:  Recent population expansion of Barred Owls (  Strix varia ) into western North America has led to concern that they may compete with and further harm the Northern Spotted Owl (  S. occidentalis caurina ), which is already listed as threatened under the U.S. Endangered Species Act (ESA). Because they hybridize, there is a legal need under the ESA for forensic identification of both species and their hybrids. We used mitochondrial control-region DNA and amplified fragment-length polymorphism (AFLP) analyses to assess maternal and biparental gene flow in this hybridization process. Mitochondrial DNA sequences (524 base pairs) indicated large divergence between Barred and Spotted Owls (13.9%). Further, the species formed two distinct clades with no signs of previous introgression. Fourteen diagnostic AFLP bands also indicated extensive divergence between the species, including markers differentiating them. Principal coordinate analyses and assignment tests clearly supported this differentiation. We found that hybrids had unique genetic combinations, including AFLP markers from both parental species, and identified known hybrids as well as potential hybrids with unclear taxonomic status. Our analyses corroborated the findings of extensive field studies that most hybrids genetically sampled resulted from crosses between female Barred Owls and male Spotted Owls. These genetic markers make it possible to clearly identify these species as well as hybrids and can now be used for research, conservation, and law enforcement. Several legal avenues may facilitate future conservation of Spotted Owls and other ESA-listed species that hybridize, including the ESA similarity-of-appearance clause (section 4[e]) and the Migratory Bird Treaty Act. The Migratory Bird Treaty Act appears to be the most useful route at this time.     

Transient dynamics of invasive competition: barred owls, spotted owls, habitat, and the demons of competition present

The recent range expansion of Barred Owls (Strix varia) into the Pacific Northwest, where the species now co-occurs with the endemic Northern Spotted Owl (Strix occidentalis caurina), resulted in a unique opportunity to investigate potential competition between two congeneric, previously allopatric species. The primary criticism of early competition research was the use of current species' distribution patterns to infer past processes; however, the recent expansion of the Barred Owl and the ability to model the processes that result in site occupancy (i.e., colonization and extinction) allowed us to address the competitive process directly rather than inferring past processes through current patterns. The purpose of our study was to determine whether Barred Owls had any negative effects on occupancy dynamics of nesting territories by Northern Spotted Owls and how these effects were influenced by habitat characteristics of Spotted Owl territories. We used single-species, multi-season occupancy models and covariates quantifying Barred Owl detections and habitat characteristics to model extinction and colonization rates of Spotted Owl pairs in southern Oregon, USA. We observed a strong, negative association between Barred Owl detections and colonization rates and a strong positive effect of Barred Owl detections on extinction rates of Spotted Owls. We observed increased extinction rates in response to decreased amounts of old forest at the territory core and higher colonization rates when old-forest habitat was less fragmented. Annual site occupancy for pairs reflected the strong effects of Barred Owls on occupancy dynamics with much lower occupancy rates predicted for territories where Barred Owls were detected. The strong Barred Owl and habitat effects on occupancy dynamics of Spotted Owls provided evidence of interference competition between the species. These effects increase the importance of conserving large amounts of contiguous, old-forest habitat to maintain Northern Spotted Owls in the landscape.